Morphology and ultrastructure of sense organs on the ovipositor of Trybliographa rapae,a parasitoid of the cabbage root fly

The ovipositor of the parasitoid wasp Trybliographa rapae was examined by scanning and transmission electron microscopy. Characteristic peg-like sensilla with a cuticular ring at the base are found at the tip of the ventral valves, where they occur in a characteristic arrangement of triplets. The unusual basal structure probably protects the sensilla against damage during movement through the substrate and piercing of the host cuticle. These sensilla are each innervated by six dendrites, some of which have lamellated tips, generally considered to be characteristic of thermosensitivity. It is suggested that the remaining dendrites are gustatory, and as such probably respond to factors present in host haemolymph. A second type of peg-like sensillum is found on both the dorsal and the ventral valves. These are set in deep pits so that only the tip of the peg protrudes above the surface of the cuticle. These occur along the length of the ovipositor shaft and ultrastructural studies reveal the pegs to be innervated by a single mechanosensitive dendrite, probably monitoring the movement of the ovipositor through the substrate.     

A new mechanosensory organ on the anterior spinnerets of the spiderCupiennius salei (Araneae,Ctenidae)

We describe hitherto unknown mechanoreceptors on the anterior spinnerets of the spiderCupiennius salei. These receptors are found at the base of the spigots of the major ampullate glands which produce the dragline used by the spider as a safety thread in various behavioral situations. There are 40–60 mechanoreceptors associated with two spigots of each anterior spinneret. They are likely to provide information on the forces pulling on the dragline and also on its orientation in space. A single sensillum consists of a hole in the cuticle covered by a thin cuticular membrane. It much resembles spider slit sensilla, which are known to detect strains in the exoskeleton. Each sensillum is supplied by two dendrites most likely belonging to two bipolar sensory cells. One of the dendrites ends at the covering membrane and the other more proximally. The sensilla are arranged with their long axes roughly parallel to the circumference of the spigots. External forces, transmitted by the dragline, result in deformation of the central part of the cuticular plate at the base of the spigots and thus in stimulation of the sensilla. This is shown electrophysiologicallly. Considering their morphology, topography, and electrophysiology, these mechanoreceptors are suggested to be important in the sensory control of dragline release by the spider.     

Ultrastructural studies of Haller's organ in the argasid tick, Argas tridentatus (Argasidae)]

Haller's organ in A. tridentatus consists of a capsule and an anterior group of sensilla. The capsule is the hollow in the cuticle on the dorsal surface of the first tarsus, where 4 pored hairs of olfactory sensilla are situated under the cover of the roof, formed by an anostomosis of the upper brunches of pleomorphs (capsule's bottom non-sensory cuticular outgrowths). The canal of the accessory ampullaceous sensillum opens in a capsule near the bottom. The anterior group of sensilla consists of two parts: proximal part, containing pored grooved and thin hairs, is homologous to the anterior grouf of ixodid ticks, and distal one which has no homologues in ixodids. Fine structure of all the sensilla in the mentioned parts of Haller's organ is described in detail.     

Ultrastructure and development of single-walled sensilla placodea and basiconica on the antennae of the Sphecoidea (Hymenoptera : aculeata)

While the pore plates of some species of the Sphecoidea (Hymenoptera) rise above the antennal surface, those of other species are flush with it. Not all species possess pore plates. On the antennae of those species, which lack pore plates, small sensilla basiconica are found. The pore plates of Psenulus concolor were studied in detail. The cuticular apparatus rises above the antennal surface. Cuticular features are the encircling ledge and delicate cuticular ledges reinforcing the perforated plate, as well as a joint-like membrane that anchors the plate into the antennal cuticle. Each pore plate is associated with 9–23 sense cells and 4 envelope cells, the second of which is doubled. In very early developmental stages, however, supernumerary envelope cells are observed; they degenerate before the cuticulin layer is secreted. Envelope cell 1 secretes a temporary dendrite sheath, while the envelope cells 2–4 are responsible for the secretion of the cuticular apparatus.The morphology and the development of the small sensilla basiconica are described in Trypoxylon attenuatum. The curved sensillum pointing to the tip of the antenna is anchored by a joint-like membrane. About 15 sense cells innervate the sensillum. The number and the arrangement of the envelope cells resemble that of the sensilla placodea. During very early developmental stages, supernumerary envelope cells are also observed. They degenerate before the cuticle of the cone is secreted by the surviving envelope cells 2–4.     

The infrared receptor of Melanophila acuminata De Geer (Coleoptera: Buprestidae): ultrastructural study of a unique insect thermoreceptor and its possible descent from a hair mechanoreceptor

The paired infrared organs of Melanophila acuminata consist of 50-100 sensilla situated at the bottom of a pit next to the coxae of the mesothoracic legs, where no exocuticle is developed. Each sensillum is accompanied by a wax gland and has a cuticular lens-like spherule (diameter 12-15 mum) bulging out with its upper hemisphere above the surface, covered only by a thin cuticle of about 1 mum. Distal processes of two enveloping cells surround the entire spherule in the form of a flattened protoplasmatic layer with the exception of a small apical stalk connecting the spherule to the outer cuticle. The spherule is innervated by a single sensory neuron of the ciliary type which is anchored ventrally with the distal tip of its cylindrical and unbranched DOS in the spherule. The insertion of the dendrite, which contains a well-developed tubular body, is always eccentric like in a hair mechanoreceptor (sensillum trichodeum) and there is no evidence of any optical function of the spherule. Three enveloping cells exist, but only one - probably the trichogen cell - forms a relatively small outer receptor lymph cavity. In the posterior wall of the pit - where exocuticle is developed - so-called suppressed systems can be found which remain completely below the cuticle with their otherwise well-developed spherules. Additionally, there is a tendency towards basally flattening and longitudinally stretching of spherules which are situated more peripherally. They strongly resemble the basal regions of hair mechanoreceptors (sensilla trichodea) in their immediate neighbourhood which are also accompanied by wax glands. Because of the existence of these transitional stages and the great ultrastructural resemblance between infrared receptors and hair mechanoreceptors concerning the bauplan of the sensory neurons and their mode of innervating the cuticular apparatus, we conclude that the infrared sensilla are probably derived from hair mechanoreceptors. Based on these results and transmission measurements of infrared radiation through the cuticular components of the organ, a model of the possible function of the infrared receptor is presented.     

The fine structure of campaniform sensilla on the halteres of Drosophila melanogaster

The campaniform sensilla on halteres of Drosophila were studied by electron microscopy in order to establish the relationships of functional elements in the sensory system. The surface of the sensillum consists of an oval cuticular cap membrane which may contain resilin, the rubberlike protein. A border of denser cuticle rings the cap membrane, and extending down around the neural process is a third type of cuticle filled with a fourth light fibrous type. The four cuticular components form a system for displacement of the neural process. The neural process is differentiated into a terminal fan-shaped structure projecting from a bulbous dilatation which tapers to a neck region ending proximally with two basal bodies. The neural process is packed with microtubules. Surrounding the dendrite is an inner enveloping cell, attached to the basal body region by septate desmosomes and by desmosomes to which microtubules of the enveloping cell are applied. An outer enveloping cell surrounds the inner one. The tip of the neural process is covered with a dense secretion which is tightly bound to the cap membrane. The dense secretion is surrounded by an extracellular fluid which might be compressed hydraulically by the cuticular system. The stimulus of cuticular distortion could thus be transmitted to the neural process which may be displaced between its fixed ends.     

Chemoreceptor sensillum structure in Limulus

The chemoreceptors of Limulus polyphemus (L.) are polyneuronal sensilla found in the spines of the coxal gnathobases of each walking leg, the spines of the chilarial appendages, and the chelae of all the limbs. Each sensillum contains 6–15 bipolar sensory cells that share a single pore in the cuticle. The dendrites of the sensory cells of each sensillum course to the cuticle together. These attenuate sharply and enter a canal in the cuticle as a very narrow terminal thread. The dendrites retain their identity in the thread, but with the light microscope, they are usually not visible individually. Each thread, consisting of 6–15 dendrites, is accompanied to the cuticular surface by a cuticular tubule found within the canal. The chemoreceptor sensilla of the gnathobase, chilarium, and chela, the temperature organs of Patten, and the flabellar receptor organs all have the same basic organization. In general this is the same structural plan shown by chemoreceptors of other arthropods. Several different mechanisms of peripheral physiological interaction among receptor cells are possible with a sensillum organization like that described here for Limulus.     

Embryonic development and molting of the antennal coeloconic no pore- and double-walled wall pore sensilla in Locusta migratoria (Insecta,Orthopteroidea)

Summary The embryonic development of antennal coeloconic sensilla was studied at four stages between 132 and 252 h after oviposition in Locusta migratoria. Initially the anlagen of the sensilla consist of 2–4 sensory cells and 3 enveloping cells. Two additional cells contribute later to the formation of socket and pit. The dendritic outer segments of the sensory cells elongate before the trichogen process grows out (ecdysis type I) with exception of one sensory cell in anlagen of poreless (np) sensilla. Other differences between np and double-walled wall pore (dw wp) sensilla are not visible until at least about 220 h after oviposition. Molting, which was studied in four stages, follows ecdysis type I in both sensillum types. The fourth enveloping cell maintains its tight connection to the socket of the sensillum even after apolysis. Its apical portion is torn off and shed together with the old cuticle. The electron-dense material between the dendritic sheath and the cuticular wall of the peg in np sensilla, which is regarded important for stimulus transmission, is not deposited during retraction of the trichogen cell. The concentric walls and spoke channels characteristic of dw wp sensilla result from deposition of cuticular material around wedge-shaped projections of the trichogen cell. The typical trilaminar 15 nm cuticulin layer is produced only on the ridges of these sensilla. The first cuticular lining of the spoke channels is only 7 nm thick and of a different structure. A flocculent material surrounds the outgrowing trichogen process. It is continuous with the filling of the spoke channels and can thus be considered as component of the stimulus-transmitting material in the functioning intermolt dw wp sensilla.     

Sensory Organs in Ips typographus (Insecta: Coleoptera) Fine structure of the sensilla of the maxillary and labial palps*

The labial and maxillary palps of the bark beetle, Ips typographus, possess a diversified array of sensilla. There are four types of sensilla possessing a single tubular body indicating a mechanoreceptive function. The variation of the associated cuticular structures of these sensillar types ranges from long bristles to cavities within the cuticle. Terminal pore sensilla with a supposed mechanosensory/gustatory function and single-walled presumably chemoreceptive sensilla are found on the apical part of the terminal palp segments. A poreless sensillar type is found on the lateral side of the terminal segment of the maxillary palp. The functional capabilities of this sensillar type are presently unknown.     

A new technique for measurement of water permeability of stomatous cuticular membranes isolated from Hedera helix leaves

Transpiration of cuticular membranes isolated from the lower stomatous surface of Hedera helix (ivy) leaves was measured using a novel approach which allowed a distinction to be made between gas phase diffusion (through stomatal pores) and solid phase diffusion (transport through the polymer matrix membrane and cuticular waxes) of water molecules. This approach is based on the principle that the diffusivity of water vapour in the gas phase can be manipulated by using different gases (helium, nitrogen, or carbon dioxide) while diffusivity of water in the solid phase is not affected. This approach allowed the flow of water across stomatal pores ('stomatal transpiration') to be calculated separately from the flow across the cuticle (cuticular transpiration) on the stomatous leaf surface. As expected, water flux across the cuticle isolated from the astomatous leaf surface was not affected by the gas composition since there are no gas-filled pores. Resistance to flux of water through the solid cuticle on the stomatous leaf surface was about 11 times lower than cuticular resistance on the astomatous leaf surface, indicating pronounced differences in barrier properties between cuticles isolated from both leaf surfaces. In order to check whether this difference in resistance was due to different barrier properties of cuticular waxes on both leaf sides, mobility of 14C-labelled 2,4-dichlorophenoxy-butyric acid 14C-2,4-DB) in reconstituted cuticular wax isolated from both leaf surfaces was measured separately. However, mobility of 14C-2,4-DB in reconstituted wax isolated from the lower leaf surface was 2.6 times lower compared with the upper leaf side. The significantly higher permeability of the ivy cuticle on the lower stomatous leaf surface compared with the astomatous surface might result from lateral heterogeneity in permeability of the cuticle covering normal epidermal cells compared with the cuticle covering the stomatal cell surface.     

Campaniform sensilla of Calliphora vicina (Insecta,Diptera)

Summary A classification scheme of campaniform sensilla using morphological criteria was developed. All variations of the two most important outer structural elements, the cuticular cap and the cuticular collar, were taken into consideration: (a) the external shape of the cuticular cap; (b) the position of the cuticular cap in relation to the remaining cuticle; (c) the position of the cuticular collar in relation to the cuticular cap. This resulted in a classification of campaniform sensilla into 24 types. This typology was applied to the campaniform sensilla of Calliphora, which show considerable variations in their outer structures. According to SEM (scanning electron microscope) pictures and TEM (transmission electron microscope) sections we found only 9 out of 24 different types of campaniform sensilla in the fly.     

Ultrastructural Analysis of the Sensilla of Austroperipatus aequabilis Reid, 1996 (Onychophora,Peripatopsidae)

The head of Austroperipatus aequabilis bears five types of sensilla. which were examined by electron microscopy. They differ from each other in position, shape of outer sensory elements and cuticular socket structures. Thus, we distinguish sensilla with sensory hairs, sensilla with sensory bulbs, cone-shaped sensilla. sensilla with sensory bristles, and sensilla of the lips. They are composed of up to 15 cells, which can he separated into four cell types. The most frequent cell type is the bipolar receptor cell that occurs in all sensilla. The apical surface of this primary receptor cell is characterized by one or two partly branched cilia with a basal 9 × 2 + 0 pattern of microtubules. A modified bipolar receptor cell was found in all sensilla bearing a sensory peg except for the sensilla equipped with sensory bristles. The apical dendrite extends to a long pale process which exclusively contains mitochondria and single microtubules. In all sensilla examined in this study at least one supporting cell occurs which is characterized by parallel microvilli. An additional function of this cell type as a part of the stimulus-conducting system is possible. In the sensillum with a sensory bulb two kinds of supporting cells occur. A unique cell type with an upside down position has regularly been found in all sensilla bearing a sensory peg. Apart from the sensilla they also occur within the labial epidermis. Since most sensilla contain several different receptor cells, they can be considered as complex sense organs. © 1998 The Royal Swedish Academy of Sciences. Published by Elsevier Science Ltd. All rights reserved     

The morphology of spider sensilla. I. Mechanoreceptors

The common tactile hair sensilla of spider tarsi were studied in web spiders (Araneus) and ground spiders (Lycosa, Dugesiella) using scanning and transmission electron microscopy. All of these sensilla are innervated by three bipolar neurons whose dendrites end proximally at the sensillum base. Each dendritic terminal exhibits a tubular body, a dense array of microtubules typical for mechanoreceptive sensilla. A dendritic sheath encloses the outer dendritic segments and connects the dendritic terminals to cuticular components of the hair sensillum in three different ways: (1) A distal extension of the dendritic sheath connects to the midline of the hair base; (2) A forked arrangement of cuticular (?) strands attaches on both lateral sides of the hair base, and (3) The socket cuticle directly contacts a part of the dendritic sheath. The latter connection provides a fixed position for the three dendritic terminals and any movement of the hair shaft could be transmitted via connections (I) and (2). The triple innervation strongly suggests a directional sensitivity of these sensilla.Structural comparison between arachnid and insect mechanoreceptive sensilla indicates that tactile hair sensilla in Arachnida are multi-innervated whereas the corresponding reccptors in Insecta are singly innervated.     

Development of the crystalliferous cuticle of Chamaecyparis lawsoniana (A. Murr.) Pari. (Gupressaceae)

A developmental study of the cuticle has shown that it consists of a homogeneous cuticle proper apposed on the wall and a heterogeneous cuticular layer generated by intussusception of cutin into the wall. At an early stage, the adcrusted cuticle proper is underlain by a ruthenium red-positive layer in which the cuticular layer originates. The origin of the anticlinal flange is referable to an electron-dense, ruthenium red-positive ridge which arises above the anticlinal wall and which also becomes cutinized. At leaf maturity, the inner surface of the cuticular layer, including that of the flange, forms interdigitating protuberances with the cell wall.
Development of the cuticle coincides with deposition of crystals of calcium oxalate in the epidermal cell wall. Initiation of large, early-formed crystals is associated with electron-opaque membranous structures formed close and parallel to the plasmalemma in the young cell wall. Crystals undergo periclinal and anticlinal growth and subsequently become engulfed within the cuticle by development of the cuticular layer. Cutin/polysaccharide interaction during development and the significance of crystal deposition are discussed.     

Ultrastructure of the antennal sensilla of aphids

Summary An electron microscopical study was made of the coeloconic and placoid sensilla on the antennae of the aphids Aphis pomi, Macrosiphum euphorbiae, Nasonovia ribis-nigri, and Pemphigus bursarius. Scanning electron microscopy revealed some variation in morphology which may be functionally important but is more likely to reflect the evolution of these species.The placoid sensilla were shown by transmission electron microscopy to have the same basic structural pattern. Each group of two or three neurons is surrounded by two ensheathing cells. The ciliary regions of the dendrites pass through a vacuole into a cavity between an outer and an inner cuticle where they may be connected to the dendritic branches although such connections were not seen. Small pores (8 nm diameter) partially penetrate the cuticle implying that these sensilla have an olfactory function. They are suggested to be important in host selection by alate aphids.The coeloconic sensilla are poreless pegs with nonsensory cuticular projections at their tips. The distal portions of their dendrites contain densely packed microtubules and the cellular arrangement of the sensilla is similar to that of the placoid sensilla. It is suggested that they may function as thermoreceptors.The authors thank the Long Ashton Research Station, Bristol for use of the SEM facilities. A.K. Bromley gratefully acknowledges the tenure of a S.R.C. CASE Studentship and thanks Professor L.H. Finlayson for research facilities     

Morphology of the prothoracic discs and associated sensilla of Acanthocnemus nigricans (Coleoptera, Acanthocnemidae)

The Australian ‘little ash beetle’ Acanthocnemus nigricans (Coleoptera, Cleroidea, Acanthocnemidae) is attracted by forest fires. A. nigricans has one pair of unique prothoracic sensory organs and it has been speculated that these organs may play a role in fire detection. Each organ consists of a cuticular disc, which is fixed over an air-filled cavity. On the outer surface of the disc, about 90 tiny cuticular sensilla are situated. The poreless outer peg of a sensillum is 3–5 μm long and is surrounded by a cuticular wall. One ciliary sensory cell innervates the peg. As a special feature, the outer dendritic segment is very short already terminating below the cuticle. A massive electron-dense cylindrical rod, which most probably represents the hypertrophied dendritic sheath, extends through the cuticular canal connecting the tip of the outer dendritic segment to the peg. The dendritic inner segment and the soma are fused indistinguishably. Thin, leaflike extensions of glial cells deeply extend into that conjoint and considerably enlarged compartment which also contains large numbers of mitochondria. In summary, the sensilla of the sensory disc of A. nigricans represent a new type of insect sensillum of hitherto unknown function. The possible role of the prothoracic sensory organ in fire detection is discussed.     

Proprioceptors involved in stinging response of the honeybee, Apis mellifera

Two types of mechanosensitive proprioceptor organ are present on the stinging apparatus of the honeybee: campaniform sensilla and mechanosensory hairplates. The campaniform sensilla are located on the surface of the tapering sting-shaft, which comprises an unpaired stylet and paired lancets. Each sensillum on the lancet differs from that on the stylet in terms of their topography and external morphology. The sensory afferents of the campaniform sensilla display slow-adapted firing responses to deformation of the cuticle that would be caused by the action of inserting the sting into a substrate, and their afferent signals induce and/or prolong the stinging response. By contrast, the mechanosensory hairplates are located at basal cuticular plates and on the posterior surface of the lancet valves. Two fields of hairplates on the second ramus at the ventral edge of the groove and on the antero-lateral edge of the oblong plate respond synchronously to protraction of the lancet. During the stinging response, these hairplates are likely to detect any sliding movement of the lancet and its position relative to the stylet. Afferent signals produced by them are likely to provide important information to the neuronal circuit for the generation and modulation of the stinging motor pattern.     

Apical antennal sensilla in nymphs of Libellula depressa (Odonata: Libellulidae)

Abstract. In an ultrastructural study of the apical antenna of the last nymphal stages of Libellula depressa (Odonata: Libellulidae), we found long sensilla trichodea, 2 sensory pegs, and a coeloconic sensillum on the last article of the flagellum (the distal part of the antenna). The long sensilla trichodea are mechanoreceptors, almost identical to the long filiform hairs of some terrestrial insects and the first sensilla of this kind to be described in aquatic insects. Particular attention was given to the complex coeloconic sensillum, a compound sensillum innervated by 2 groups of 3 neurons wrapped in a dendritic sheath. A cuticular sleeve envelops the distal portion of the outer dendritic segment. The cuticle of the coeloconic sensillum shows wide channels and is contiguous to the underlying granular and fibrillar layer. Similar structures on the antennae of the adults of other dragonflies were identified as chemoreceptors in previous studies. We hypothesize that this larval coeloconic sensillum might likewise have a chemosensory function, responding to molecules that diffuse through the cuticle and the underlying granular and fibrillar layer, as no clear pore or pore-tubule system is visible. Alternative functions are also explored on the basis of morphological details.     

Structure and localization of sensilla on antennas of caddisflies (insecta: trichoptera)

The structures of antennal segments and ultrastructures of antennal sensilla were studied in representatives of 28 families of caddisflies from both extant suborders by the methods of light and scanning electron microscopy. Sixteen types of the sensilla have been found to occur on the antenna in Trichoptera; some of them were found for the first time. Morphological characters of the cuticular structures on the antennal surface demonstrate the significant structural differences both in various families and in the lower taxonomy levels. Specialized sensory fields differing structurally from the rest of the flagellomer surface have been found on the antennas in the suborder Phryganeina. A modified classification of sensilla based on the cuticular structures is proposed.